Pinheiro et al 2009 Zootaxa by Voz da Natureza

Zootaxa 2298: 45–54 (2009) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Article

ZOOTAXA ISSN 1175-5334 (online edition)

New records of fishes for Trindade-Martin Vaz oceanic insular complex, Brazil HUDSON T. PINHEIRO, VICTOR CAMILATO, JOÃO LUIZ GASPARINI & JEAN-CHRISTOPHE JOYEUX Departamento de Oceanografia e Ecologia, Universidade Federal do Espírito Santo, Av. Fernando Ferrari, 514, Goiabeiras, 29075910, Vitória, ES, Brazil. E-mail: htpinheiro@gmail.com; vcamilato@yahoo.com.br; gaspa.vix@terra.com.br; joyeux@npd.ufes.br

Abstract Thirty-two new records for shore fishes were made at the Trindade-Martin Vaz oceanic insular complex, located 1,160 km off the Brazilian coast. These records are related to an increase in sampling effort and to temporal variation in population size. Newly found but very abundant species hypothetically exemplify temporal variations in population size. The Vitória-Trindade submarine chain also may function as a series of stepping-stones facilitating transport of typically coastal species. The new records reveal a lack of knowledge of the fish fauna of this unique location. Studies aimed at the exploration of shallow areas, deep reefs and seamounts of the Vitória-Trindade chain, aided by ichthyoplanktonic surveys, are essential to a better understanding of the fish community and the processes of colonization in this oceanic locality. Key words: South Atlantic; distribution; biogeography; Brazil

Introduction The colonization and maintenance of marine populations at remote islands are important and much discussed themes (Gad & Schminke 2002), but these areas still remain poorly explored (Forges et al. 2000). Processes of colonization of oceanic islands can involve dispersal aided by submarine mounts acting as stepping-stones (Edwards 1993; Moore et al. 2004), whereas the maintenance of species at islands and seamounts is usually attributed to oceanic processes stimulating larval retention (Mullineaux & Mills 1997). Even so, community composition at isolated oceanic islands reportedly experiences large spatial and temporal variations (Stobberup et al. 2002). Even though there often is a large ratio of endemics at remote islands (that may reach over 30% - Forges et al. 2000), little is known about the actual distribution of these species considering that they may also occur in 'nearby' unexplored seamounts (Edwards 1993). Trindade Island (20°31′ S, 29°19′ W) and Martin Vaz Archipelago (20°30′ S, 28°51′ W), located about 1,160 km off the coast of Brazil, are the only emerged components of an sunken submerged volcanic chain composed by eight mountains disposed perpendicularly to the coast of the state of Espírito Santo, southeastern Brazil. The mountains of the Vitória-Trindade chain reach between 10 and 110 meters below the water surface and are separated by about 250 km from each other (Figure 1). Several biogeographic studies have included ichthyofaunal data from Trindade (e.g., Floeter & Gasparini 2000; Floeter et al. 2001; Joyeux et al. 2001; Floeter et al. 2008), and outlined similarities and contrasts with the Brazilian coastal ichthyofauna. However, in contrast to the other Brazilian and Atlantic Central oceanic islands (St. Paul´s Rocks, Fernando de Noronha, Atol das Rocas, St. Helena and Ascension – Lubbock 1980; Lubbock & Edwards 1981; Edwards & Lubbock 1983; Edwards & Glass 1987a, b; Rosa & Moura 1997; Soto 2001; Feitoza et al. 2003; Sampaio et al. 2006), baseline data for Trindade-Martin Vaz ichthyofauna remains poor. Only one recent ichthyological expedition was conducted, which identified 97 species, including six

Accepted by L. Rocha: 29 Oct. 2009; published: 26 Nov. 2009

endemic species and one sub-species (Gasparini & Floeter 2001). Similarly to what happens at other Brazilian oceanic islands (Debelius 1997; Oliveira et al. 1997), fisheries represent the foremost human influence to the Trindade-Martin-Vaz ichthyofauna. In three recent scientific expeditions, we recorded the presence of 32 fish species previously undocumented at the insular complex. This work presents the new records and discusses potential factors that may have led to these species being overlooked previously.

FIGURE 1. Profile of the Vitória-Trindade chain and the Trindade-Martin Vaz insular complex located off the central coast of Brazil (T= Trindade island; MV= Martin Vaz archipelago). Modified from Almeida (2006).

Material and methods Study area: Trindade Island and Martin Vaz Archipelago are located over two different submerged mountains that have their bases at 4000 meters depth. Trindade Island has 9.3 km² of land area and about 32 km² of shallow shelf (0-50 m; Gasparini & Floeter 2001). Calcareous algae, volcanic rocks and sandy plateau are the main components of a substrate that is poorly colonized by few species of corals, zoanthids and sponges. Martin Vaz is composed by three small islands located 47 km east of Trindade and is Brazil’s most distant offshore locality. Submerged substrate is characterized by high complexity arising from large rocks covered by calcareous algae. Sampling and data analysis: Ichthyofaunal records result from 212 hours of observation (130 dives) at Trindade and 15 h at Martin Vaz spread over three periods: January-February and July-September 2007 and April-June 2009. Records consist of field identifications and photographs. A number of individuals were collected by hand-net or spear-gun. These individuals were deposited in the ichthyological collection of the Universidade Federal do Espírito Santo (CIUFES). The habitat characterization follows Gasparini & Floeter (2001) and the geographic range was attributed using Floeter et al. (2008). Families are listed in phylogenetic order following Nelson (2006) and alphabetic order within families. Identifications were made using Figueiredo (1977), Figueiredo and Menezes (1978; 1980; 2000), and Menezes and Figueiredo (1980; 1985).

Results Thirty-two new records of fishes in 23 families are documented for the Trindade-Martin Vaz oceanic insular complex (Table 1). Families with the most new members are Labridae (n = 3), Apogonidae, Carangidae, Diodontidae, Holocentridae, Pempheridae, Scaridae and Tetraodontidae (n = 2). Fifteen new records are of demersal species, eight benthonic, seven pelagic and two are of mid-water species. In respect to zoogeographic distribution patterns, eleven are distributed throughout the Western Atlantic, nine are panAtlantic, three occur on both sides of the Atlantic (west and east), three are endemic to Trindade and Martin Vaz, two are distributed in both the Mid-Atlantic Ridge and the Western Atlantic, one has been reported from

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FIGURE 2. Examples of reef fishes recorded for the first time at the Trindade-Martin Vaz oceanic insular complex. Aetobatus narinari (a); Sargocentron bullisi (b); Scorpaenodes caribbaeus (c); Cephalopholis furcifer (d); Alectis ciliaris (e); Caranx crysos (f); Pempheris poeyi (g); Amblycirrhitus pinos (h). Photos by V. Camilato (a, d, e); H. Pinheiro (b, f); R. Santos (c, h); J. Gasparini (g). NEW RECORDS FOR TRINDADE - MARTIN VAZ

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FIGURE 3. Examples of reef fishes recorded for the first time at the Trindade-Martin Vaz oceanic insular complex. Halichoeres sp. (a); Cryptotomus roseus (b); Hypleurochilus sp. (c); Canthigaster figueiredoi (d); Sphoeroides spengleri (e); Chilomycterus reticulatus (f); Cyclichthys spinosus (g); Hybrid between C. furcifer and C. fulva (h). Photos by R. Santos (d, f); H. Pinheiro (the others).

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TABLE 1. New records for Trindade-Martin Vaz reef fishes. Habitat, depth, abundance (number of individuals per number of dives found), geographic range, sampling method, site (detailed for Trindade) and voucher are showed. (The taxa Hypleurochilus sp. is not technically a new record as it has been reported by Gasparini & Floeter (2001) as H. fissicornis.) Family / Species

Habitat

Depth (m)

Abundance (ind. / dives)

Geographic Range

Sample Method

Site

Voucher

1/1

Praia do Andrada

1/2

Ponta da Calheta; Praia Figure 2, a dos Cabritos

0 - 10

3/3

Ponta da Calheta; Praia do Lixo, das Tartarugas

0 – 0.5

10 / rockshore

End.

Praia das Tartarugas, dos Portugueses

CIUFES 1492

Plectrypops retrospinis (Guichenot, 1853)

1/1

Ponta da Calheta

Sargocentron bullisi (Woods, 1955)

9 - 25

3/2

Parcel; Ponta da Calheta

Figure 2, b

1/1

Praia do Andrada

Figure 2, c

5 - 17

14 / 3

Parcel; Praias do Andrada, da Calheta, do Noroeste and das Tartarugas

Figure 2, d

1/1

SWA+MAR+ MC EA

Martin Vaz

CIUFES 0849

6-8

6/2

Ponta da Calheta

CIUFES 0856

Astrapogon stellatus (Cope, 1867)

1-2

5/1

Praia do Lixo

CIUFES 0845

Phaeoptyx pigmentaria (Poey, 1860)

2-3

6/2

WA+EA

Ponta da Calheta

CIUFES 0844

1/1

Parcel

Alectis ciliaris (Bloch, 1787)

2/1

WA+EA

Pedra da Garoupa; Ponta da Calheta

Figure 2, e

Caranx crysos (Mitchill, 1815)

3/1

Martin Vaz; Ponta da Calheta

Figure 2, f

0-1

12 / 1

NWA+TR

Poça do Parcel

CIUFES 1493; Figure 2, g

Rhincodontidae Rhincodon typus Smith, 1828 Carcharhinidae Galeocerdo cuvier (Péron & Lesueur, 1822) Myliobatidae Aetobatus narinari (Euphrasen, 1790) Muranidae Enchelycore carychroa Böhlke & Böhlke, 1976 Gobiesocidae Tomicodon sp. Holocentridae

Scorpaenidae Scorpaenodes caribbaeus Meek & Hildebrand, 1928 Serranidae Cephalopholis furcifer (Valenciennes, 1828)

Aulostomidae Aulostomus strigosus Wheeler, 1955 Dactylopteridae Dactylopterus volitans (Linnaeus, 1758) Apogonidae

Echeneidae Remora remora (Linnaeus, 1758) Carangidae

Pempheridae Pempheris poeyi Bean, 1885

continued next page

NEW RECORDS FOR TRINDADE - MARTIN VAZ

TABLE 1. (continued) Family / Species

Habitat

Depth (m)

Abundance (ind. / dives)

Geographic Range

Sample Method

Site

Voucher

Pempheris schomburgkii Müller & Troschel, 1848

3-6

15 / 2

Praia das Cabritas

CIUFES 0850

1/1

WA+MAR

Ponta da Calheta

Figure 2, h

10 - 15

4/2

Ponta da Calheta

CIUFES 0857

Clepticus brasiliensis Heiser, Moura & Robertson, 2000

3/1

Ponta da Calheta

CIUFES 0851

Halichoeres sp.

6 - 25

>50 / 5

End.

Martin Vaz; Ponta da Calheta

CIUFES 0858; Figure 3, a

Xyrichtys splendens (Castelnau, 1855)

4/2

Ponta da Calheta

CIUFES 1008

Cryptotomus roseus Cope, 1871

2/1

WA+MAR

Ponta da Calheta

CIUFES 1494; Figure 3, b

Sparisoma sp.

5 - 12

5/3

End.

Parcel; Praias do Noroeste, das Tartarugas

CIUFES 0848

4 - 12

25 / 15

End.

Ponta da Calheta

Figure 3, c

12 - 18

8/5

Enseada do Farol

CIUFES 1495

6 - 40

5/2

Ponta da Calheta; Praia CIUFES das Cabritas, do Tunel 1496

5/3

Praia das Cabritas; Calheta

10 - 20

10 / 10

Praia do Lixo; Calheta

Figure 3, d

1/1

WA+EA

Praia do Lixo

Figure 3, e

Chilomycterus reticulatus (Linnaeus, 1758)

10 - 20

10 / 10

Ponta da Calheta

Figure 3, f

Cyclichthys spinosus (Linnaeus, 1758)

10 -2 0

4/4

SWA

Ponta da Calheta

Figure 3, g

Cirrhitidae Amblycirrhitus pinos (Mowbray, 1927) Pomacentridae Stegastes pictus (Castelnau, 1855) Labridae

Scaridae

Blenniidae Hypleurochilus sp. Callionymidae Callionymus bairdi Jordan, 1888 Gobiidae Coryphopterus thrix Böhlke & Robins, 1960. Scombridae Thunnus obesus (Lowe, 1839) Tetraodontidae Canthigaster figueiredoi Moura & Castro, 2002 Sphoeroides spengleri (Bloch, 1785) Diodontidae

Habitat: D= demersal; B= benthonic; P= pelagic; MW= mid-water. Geographic Range: WA= West Atlantic; SWA= Southwestern Atlantic; NWA= Northwestern Atlantic; EA= Eastern Atlantic; MAR= Middle Atlantic Ridge; BR= Brazilian province; PA= pan-Atlantic (occur in all provinces); TR= Trindade; End.= Endemic to Trindade. Sampling Method: VR= visual record; PR= photographic record; MC= manual collection.

the Southwestern, Eastern and Middle Atlantic Ridge, one is restricted to the Southwestern Atlantic, one is endemic to the Brazilian province and one, Pempheris poeyi (Figure 2, g), has an interesting disjunct distribution as it is only known from the Northwestern Atlantic and Trindade island. Among the new records, Cephalopolis furcifer (sensu Craig & Hastings 2007) (Figure 2, d) was the most abundant and widely distributed species in the complex during the two census periods (the balistid Melichthys

PINHEIRO ET AL.

niger was the most abundant species overall). Hybrids between Cephalopholis furcifer and C. fulva (Linnaeus, 1758) were recorded (Figure 3, h) in the Parcel region at 4 - 8 m of depth. These individuals have demersal habits and morphology similar to that of C. fulva, but have a duskier coloration, deeper body and bifurcate caudal fin. Genetic studies (L. Rocha, comm. pers.) revealed that Halichoeres sp. (Figure 3, a) is a new species presently being described by the first author and colleagues. Common, this wrasse lives alone or in small aggregations (up to 15 individuals) between 6 m and at least 25 m, mostly in high complexity habitats. The parrotfish Sparisoma sp. and the clingfish Tomicodon sp. also are new species awaiting genetic studies to reveal their exact phylogenetic position. The combtooth blenny Hypleurochilus sp. (Figure 3, c), albeit not technically a new record since it is listed as H. fissicornis Quoy & Gaimard, 1824 in Gasparini & Floeter (2001), is worth mentioning since the capture of a higher number of individuals revealed this species is actually new to science. A few records are unusual for diverse reasons. Aetobatus narinari (Figure 2, a) and Cryptotomus roseus (Figure 3, b) were sighted twice while only one specimen of Aulostomus strigosus, Plectrypops retrospinis and Remora remora were observed. Galeocerdo cuvier and Rhincodon typus were recorded from recent Brazilian Navy archives.

Discussion Taxonomic comments Several endemic species of the Brazilian province have been taxonomically revised and are now validated as Halichoeres penrosei (previously recorded as H. maculipinna – review in Rocha 2004), Sparisoma tuiupiranga, S. axillare and S. amplum (previously recorded as S. atomarium, S. rubripinne and S. viride respectively in Gasparini & Floeter 2001 – reviews in Moura et al. 2002 and Gasparini et al. 2003). Two Trindade endemics, the combtooth blenny Scartella poiti Rangel, Guimarães and Gasparini, 2004 and the cleaning goby Elacatinus pridisi Guimarães, Gasparini and Rocha, 2004 have been recently described. A third endemic, Stegastes trindadensis Gasparini, Moura & Sazima, 1999 is considered to be, at most, a sub-species of Stegastes fuscus (Cuvier 1830). Descriptions of two other shallow-water species are under review. Hybrid specimens between C. fulva and C. furcifer have been observed in other Brazilian oceanic islands (Fernando de Noronha and Atol das Rocas; J.L. Gasparini & H.T. Pinheiro, pers. obs.) and in the North Atlantic (Cuba and Bermuda; Smith 1966; Bostrom et al. 2002). Such hybrids were initially described by Poey in 1860 and 1875 as Menephorus dubius and M. punctiferus (Smith 1966; Bostrom et al. 2002). Hybridization in the marine environment can occur due to the difficulty to achieve reproduction for the less abundant species (Lara-Ruiz et al. 2006), in this case C. furcifer, since C. fulva is a common species on Trindade reefs (Gasparini & Floeter 2001; pers. obs.). Difficulty of reproduction, possibly combined with a decrease in the rate of fertilization (Petersen & Warner 2002), could cause large fluctuations in the size of the population over time and such isolated populations could crash, recover or become extinct without notice. Zoogeography The 32 new records represent an increase of 33% in the checklist of Trindade-Martin Vaz complex, raising it from 97 to 129 species. The family Carangidae is the most speciose (n = 11), followed by Labridae and Serranidae (n = 10) and Muraenidae and Pomacentridae (n = 6). Taking into account that more than 70% of endemic species of the Brazilian province have demersal spawning (Floeter & Gasparini 2000), Apogonidae, Pomacentridae and Gobiidae should receive a stronger emphasis in taxonomic, genetic and phylogeographic studies. Some new records represent significant range extensions. Pempheris poeyi is registered for the first time in the South Atlantic while Sargocentron bullisi, Scorpaenodes caribbaeus, Xyrichthys splendens, Coryphopterus thrix and Cyclichthys spinosus are registered for the first time in a south Atlantic oceanic island (including northern hemisphere´s St Paul´s rock). While absent from St Paul´s Rocks, NEW RECORDS FOR TRINDADE - MARTIN VAZ

Fernando de Noronha and Atol da Rocas (the other Brazilian oceanic islands), Callionymus bairdi is also found at St Helena (but not Ascension) and Chilomycterus reticulatus at both St Helena and Ascension. Geographic distributions of P. poeyi, Amblycirrhitus pinos and Sphoeroides spengleri are unusual, the first for being highly disjunct (e.g., Joyeux et al. 2001), the second for encompassing St Helena but not Ascension (where A. pinos is replaced by its sister species Amblycirrhitus earnshawi Lubbock, 1978; Floeter et al. 2008) and the third for having a Western Atlantic distribution extending to the Canaries in the Eastern Atlantic (Floeter et al. 2008). These records are, however, consistent with what we know of Atlantic reef fish biogeography (e.g., Floeter et al. 2008), in which Trindade and Martin Vaz are impoverished outposts of the Brazilian province. Species endemic to the Trindade insular complex do appear to offer support to this close relationship with the Brazilian coast. Many sister species are SWA endemics (Elacatinus figaro Sazima, Moura & Rosa 1997 for E. pridisi; Sparisoma frondosum (Agassiz, 1831) for Sparisoma sp.; Malacoctenus aff. triangulatus for Malacotenus sp. [listed in Gasparini & Floeter 2001]; Entomacrodus vomerinus (Valenciennes, 1836) for Entomacrodus sp. [listed in Gasparini & Floeter 2001]). However, others are either pan-Atlantic (Scartella cristata (Linnaeus, 1758) for S. poiti) or with uncertain affinities (Halichoeres sp., Tomicodon sp. and Hypleurochilus sp.). The new records for reef fish species at Trindade-Martin Vaz can be attributed, primarily, to three factors. The first is the increase in sampling effort and the use of SCUBA equipment. The paucity of ichthyological studies is due to the distance between the island and the coast and the complicated logistics (including the difficulty of access to beaches from land), the large size of the shallow platform and the lack of infrastructure to support SCUBA diving. The second factor is temporal variation in population sizes, which may reduce the odds of recording such species during periods of low abundance. The third factor could be the re-colonization or colonization of the insular complex by 'new' species. This re-colonization or colonization of species is likely to be facilitated by the influence of the Vitória-Trindade seamount chain, which can act as a series of "stepping stones". Floeter and Gasparini (2000) suggested that the chain is responsible for the great affinity of the fish fauna of Trindade Island with that of the Brazilian coast. Such an affinity implies that genetic fluxes are relatively constant (relative to the necessary time for allopatric divergence) and that they would, in most cases, greatly limit the extent and frequency of population crashes. The new records presented here indicate a gap in our knowledge of the fish fauna of the Trindade-Martin Vaz island complex. In this context, future additional records are probable. Studies aimed at the exploration of deep reefs and seamounts of the Vitória-Trindade chain, basically inaccessible until now, aided by ichthyoplanktonic surveys, are essential to a better understanding of the fish community and the processes of (re)settlement in this singular oceanic locality.

Acknowledgements We thank TAMAR / ICMbio project team leaders Luciana Magnabosco, Soraya Bruno and Antonio de Padua Almeida and the researchers D. Krise, L. Bugoni, R. Santos, T. Simon, H. Filgueiras, C.E. Stein, P. Welff and L. Rocha. The logistic support of the Brazilian Navy was fundamental to the realization of this study and in particular we would like to express our gratitude to Carlos Eduardo, André, Carlos, R. Pereira, A. Lopes, W. Fernandes, P. Reis, W. da Silva, E. Souza, Ronaldo, Augusto Cândido, Luberiaga, Baruf, Freitas, Cardoso, F.L. Ruiz, J.C.G. Pereira, Rodrigues, Diego and Robson. Fundamental partnership and logistical support to diving activities was provided by diving operators Flamar and Windive and by NGO Voz da Natureza.

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